氮磷添加对色季拉山高寒草地土壤酶活性和化学计量比的影响

doi:10.16258/j.cnki.1674-5906.2026.04.003

生态环境学报 ›› 2026, Vol. 35 ›› Issue (4): 520-528.DOI: 10.16258/j.cnki.1674-5906.2026.04.003

氮磷添加对色季拉山高寒草地土壤酶活性和化学计量比的影响

崔志攀(), 张绍兵, 张西哲, 韩艳英, 吴正, 仝令臣, 叶彦辉^*()

西藏农牧大学林草学院，西藏林芝 860000

收稿日期:2025-10-08 修回日期:2026-02-03 接受日期:2026-03-02 出版日期:2026-04-18 发布日期:2026-04-14
通讯作者: *E-mail: 260383872@qq.com
作者简介:崔志攀（1997年生），男，硕士研究生，主要从事高寒草地氮磷沉降研究。E-mail: 1723957643@qq.com
基金资助:
西藏农牧学院研究生教育创新计划项目(YJS2024-31);西藏农牧学院研究生教育创新计划项目(YJS2024-26);西藏农牧学院研究生教育创新计划项目(YJS2024-28);西藏农牧学院林学博士点（一期）(533325001);国家自然科学基金项目(31860141);国家自然科学基金项目(31360119);西藏自治区科技重大专项(XZ202201ZD0005G02);西藏林芝苹果科技小院(XY2024-03);西藏农牧学院重点学科建设项目(XK20245-04);林芝市科技计划项目(2023-XYQ-006);西藏自治区科技计划重点研发计划(XZ202401ZY0032);西藏自治区科技计划重大专项(XZ202101ZD003N);2025年中央财政支持地方高校发展改革专项资金(XK2024-04);2025年中央财政支持地方高校发展改革专项资金(XK2024-01);2025年中央财政支持地方高校发展改革专项资金(YJSXK2025-22);西藏农牧学院第七批柔性人才项目(53013001804)

Effects of Nitrogen and Phosphorus Addition on Soil Enzyme Activity and Stoichiometric Ratios in the Alpine Grassland of Sejila Mountain

CUI Zhipan(), ZHANG Shaobing, ZHANG Xizhe, HAN Yanying, WU Zheng, TONG Lingchen, YE Yanhui^*()

College of Forestry and Grassland, Xizang Agricultural and Animal Husbandry University, Nyingchi 860000, P. R. China

Received:2025-10-08 Revised:2026-02-03 Accepted:2026-03-02 Online:2026-04-18 Published:2026-04-14

摘要/Abstract

摘要：

由于人类活动和工业发展活性氮磷的输入持续增多。目前，色季拉山高寒草地还缺乏氮磷富集下土壤理化性质的研究，且较少关注氮磷的交互作用。以色季拉山高寒草地为对象，通过开展野外氮磷添加控制实验，探究土壤酶活性和化学计量比对养分添加的响应机制。结果显示，1）相较对照，氮磷复合添加下β-1,4-葡萄糖苷酶与纤维二糖水解酶分别上升71.3%与62.0%（p<0.05）；氮获取酶呈“异向”响应：β-1,4-N-乙酰基氨基葡萄糖苷酶在磷与氮磷下显著升高90.6%、59.8%，在氮处理下降43.8%；亮氨酸氨基肽酶仅在氮磷复合处理下显著升高111%；碱性磷酸酶主要在氮磷复合处理下上升55.2%（p<0.05）。2）酶化学计量与矢量分析表明，微生物总体受磷限制：各处理矢量角均大于45°；氮磷虽提升群落代谢上限，但主导限制仍偏向磷端。3）相关分析与冗余分析显示，外源氮、磷及氮磷通过改变土壤碳氮比、有效磷、微生物量碳与有机碳供给等因子共同调控酶与酶计量结构；磷与氮磷下有效磷显著提高103%、427%，微生物量碳在氮磷处理下降低了32.9%，溶解性有机碳氮磷处理下降低8.21%，样点沿“有效磷－碳氮比”梯度清晰分离（p<0.05）。综上所述，该研究揭示了高寒草地在氮、磷富集下的“计量再平衡－代谢阈值”特征：单因子添加改变特定位点的“酶学投资”，复合添加放大代谢潜力但仍未改变磷主导限制。

关键词: 氮磷添加, 土壤酶活性, 酶化学计量比, 高寒草地, 色季拉山

Abstract:

Due to intensified human activities and industrial development, the input of reactive nitrogen (N) and phosphorus (P) has been continuously increasing. However, there is a lack of studies on soil physicochemical properties under N and P enrichment in the alpine grassland of Sejila Mountain, and little attention has been paid to the interactive effects of N and P. This study examined the response mechanisms of soil enzyme activities and stoichiometric ratios to nutrient addition using a field-control experiment of N and P addition in the alpine grassland of Sejila Mountain. Results showed that: 1) compared with the control, the activities of β-1,4-glucosidase and cellobiohydrolase increased by 71.3% and 62.0%, respectively, under combined N and P addition (p<0.05). Nitrogen-acquiring enzymes displayed heterogeneous responses: β-1,4-N-acetylglucosaminidase increased by 90.6% and 59.8% under P addition and combined N-P addition, respectively, but decreased by 43.8% under N addition. Leucine aminopeptidase increased by 111.2% only under combined N-P addition. Alkaline phosphatase increased by 55.2% under combined N-P addition (p<0.05). 2) Enzyme stoichiometry and vector analysis showed that the soil microorganisms were predominantly limited by P, as the vector angles under all treatments exceeded 45°. Although N and P addition elevated the metabolic upper limit of the microbial community, the dominant limitation still tended to be P. 3) Correlation analysis and redundancy analysis indicated that exogenous N, P, and their combined addition jointly regulated soil enzyme activities and their stoichiometric structure by altering soil C꞉N ratio, available phosphorus, microbial biomass carbon, and dissolved organic carbon. Under P addition and combined N-P addition, available phosphorus increased markedly by 103.1% and 427.2%, respectively; microbial biomass carbon declined by 32.9% under combined N-P addition; and dissolved organic carbon decreased by 8.2% under combined N-P addition. The sampling sites were distinctly separated along the gradient of “available phosphorus-carbon-nitrogen ratio” (p<0.05). In summary, this study demonstrates the characteristics of “stoichiometric rebalancing-metabolic threshold” of alpine grasslands under N and P enrichment: single-factor addition modifies enzymatic investment at specific sites, whereas combined addition enhances the overall metabolic potential without shifting P from its role as the dominant limiting nutrient.

Key words: nitrogen and phosphorus addition, soil enzyme activity, enzyme stoichiometry ratio, alpine grassland, Sejila Mountain

中图分类号:

崔志攀, 张绍兵, 张西哲, 韩艳英, 吴正, 仝令臣, 叶彦辉. 氮磷添加对色季拉山高寒草地土壤酶活性和化学计量比的影响[J]. 生态环境学报, 2026, 35(4): 520-528.

CUI Zhipan, ZHANG Shaobing, ZHANG Xizhe, HAN Yanying, WU Zheng, TONG Lingchen, YE Yanhui. Effects of Nitrogen and Phosphorus Addition on Soil Enzyme Activity and Stoichiometric Ratios in the Alpine Grassland of Sejila Mountain[J]. Ecology and Environmental Sciences, 2026, 35(4): 520-528.

图/表 6

图1 样地示意图审图号为GS（2024）0650号，源于国家基础地理信息中心

Figure 1 Location of sampling sites

表1 氮磷添加对高寒草地土壤理化性质的影响

Table 1 Effects on N and P additions on the physice-chemical properties of alpine grassland soils

参数	对照	氮添加	磷添加	氮磷复合添加
pH	5.83±0.01a	5.39±0.01d	5.43±0.01c	5.66±0.01b
容重/(g‧cm⁻³)	0.76±0.01b	0.74±0.02b	0.75±0.03b	1.09±0.05a
w_(有机碳)/(g‧kg⁻¹)	49.78±2.82b	67.09±4.38a	71.00±3.67a	46.16±0.97b
w_(总氮)/(g‧kg⁻¹)	4.11±0.27b	5.53±0.48a	5.64±0.33a	4.20±0.09b
w_(总磷)/(mg‧kg⁻¹)	0.81±0.03c	0.91±0.01b	1.07±0.01a	0.90±0.02b
碳氮比	12.14±0.13a	12.20±0.40a	12.59±0.09a	11.00±0.16b
碳磷比	61.92±4.26ab	73.63±4.65a	66.28±2.95a	51.67±2.03b
氮磷比	5.11±0.37ab	6.07±0.55a	5.27±0.27ab	4.70±0.15b

图2 氮、磷添加对高寒草地土壤养分和微生物量的影响 CK：对照组：氮添加；P：磷添加；NP：氮磷复合添加；不同小写字母表示处理间差异显著（p<0.05），样本重复数n=3。下同

Figure 2 Effects of N and P addition on soil nutrients and microbial biomass in alpine grassland

图3 氮、磷添加对高寒草地土壤酶活性的影响

Figure 3 Effects of N and P addition on soi enzyme activities in alpine grassland

图4 氮、磷添加对高寒草地土壤酶计量和矢量特征的影响

Figure 4 Effects of N and P additions on enzyme stoichiometric ratios and vector characteristics of alpine grassland soils

图5 土壤酶活性和酶计量比与理化因子之间的关系 BD：容重；SOM：土壤有机碳；TP：总磷；TN：总氮；C?N?碳氮比；C?P?碳磷比；N?P?氮磷比；AP：有效磷；AN：有效氮；DOC：可溶性有机碳；CMB：微生物碳；NMB：微生物氮；PMB：微生物磷；VA：矢量角度；VL：矢量长度；CAE：碳获取酶；NAE：氮获取酶；PAE：磷获取酶；Ecn：酶碳氮比；Ecp：酶碳磷比；Enp：酶氮磷比；BG：β-1,4葡萄糖苷酶；CBH：纤维二糖水解酶；NAG：B-1,4N-乙酰氨基葡萄糖苷酶；LAP：亮氨酸氨基肽酶；ALP：碱性磷酸酶；CK：对照组；N：氮添加；P：磷添加；NP：氮磷复合添加，样本重复数n=3

Figure 5 Soil enzyme activities and enzyme stoichiometric ratios in relation to physico-chemical factors

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氮磷添加对色季拉山高寒草地土壤酶活性和化学计量比的影响

Effects of Nitrogen and Phosphorus Addition on Soil Enzyme Activity and Stoichiometric Ratios in the Alpine Grassland of Sejila Mountain

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